433 lines
55 KiB
XML
433 lines
55 KiB
XML
<document ID-DOI="http://dx.doi.org/10.3897/zookeys.662.12124" ID-GBIF-Dataset="85e41c76-28ec-4f0d-86cc-6a6ffd5f3784" ID-PMC="PMC5539362" ID-Pensoft-Pub="1313-2970-662-101" ID-PubMed="28769612" ID-ZBK="0250ADB4740A4DB283FA92E3BA0363D2" ModsDocAuthor="" ModsDocDate="2017" ModsDocID="1313-2970-662-101" ModsDocOrigin="ZooKeys 662" ModsDocTitle="The ground beetle genus Bembidion Latreille in Baltic amber: Review of preserved specimens and first 3D reconstruction of endophallic structures using X-ray microscopy (Coleoptera, Carabidae, Bembidiini)" checkinTime="1490252562655" checkinUser="pensoft" docAuthor="Schmidt, Joachim & Michalik, Peter" docDate="2017" docId="ED67963CB9BF49B1A7095C5F81716414" docLanguage="en" docName="ZooKeys 662: 101-126" docOrigin="ZooKeys 662" docSource="http://dx.doi.org/10.3897/zookeys.662.12124" docTitle="Bembidion bukejsi" docType="treatment" docVersion="4" lastPageNumber="106" masterDocId="4035B67CBA16C000FFD2B459FFD5FF83" masterDocTitle="The ground beetle genus Bembidion Latreille in Baltic amber: Review of preserved specimens and first 3 D reconstruction of endophallic structures using X-ray microscopy (Coleoptera, Carabidae, Bembidiini)" masterLastPageNumber="126" masterPageNumber="101" pageNumber="104" updateTime="1668164303151" updateUser="ExternalLinkService">
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<mods:titleInfo>
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<mods:title>The ground beetle genus Bembidion Latreille in Baltic amber: Review of preserved specimens and first 3 D reconstruction of endophallic structures using X-ray microscopy (Coleoptera, Carabidae, Bembidiini)</mods:title>
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</mods:titleInfo>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Schmidt, Joachim</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Michalik, Peter</mods:namePart>
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<mods:typeOfResource>text</mods:typeOfResource>
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<mods:title>ZooKeys</mods:title>
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</mods:titleInfo>
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<mods:part>
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<mods:date>2017</mods:date>
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<mods:detail type="volume">
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<mods:number>662</mods:number>
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</mods:detail>
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<mods:extent unit="page">
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<mods:start>101</mods:start>
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<mods:end>126</mods:end>
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</mods:extent>
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<mods:location>
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<mods:url>http://dx.doi.org/10.3897/zookeys.662.12124</mods:url>
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</mods:location>
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<mods:classification>journal article</mods:classification>
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<mods:identifier type="DOI">http://dx.doi.org/10.3897/zookeys.662.12124</mods:identifier>
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<mods:identifier type="Pensoft-Pub">1313-2970-662-101</mods:identifier>
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<mods:identifier type="ZBK">0250ADB4740A4DB283FA92E3BA0363D2</mods:identifier>
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<mods:identifier type="ZooBank">0250ADB4740A4DB283FA92E3BA0363D2</mods:identifier>
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</mods:mods>
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<treatment ID-GBIF-Taxon="127879722" LSID="urn:lsid:plazi:treatment:ED67963CB9BF49B1A7095C5F81716414" httpUri="http://treatment.plazi.org/id/ED67963CB9BF49B1A7095C5F81716414" lastPageId="5" lastPageNumber="106" pageId="3" pageNumber="104">
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<subSubSection pageId="3" pageNumber="104" type="nomenclature">
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<paragraph pageId="3" pageNumber="104">
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<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion bukejsi" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="species" species="bukejsi">Bembidion bukejsi</taxonomicName>
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http://zoobank.org/661D694B-E65E-4D6F-A0D6-A26C0BC3D50EFigs 1-2, 3-5, 6-8, 9-13, 14-17, 26, 27
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</paragraph>
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</subSubSection>
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<subSubSection pageId="3" pageNumber="104" type="holotype">
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<paragraph pageId="3" pageNumber="104">Holotype.</paragraph>
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<paragraph pageId="3" pageNumber="104">
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Male in Baltic amber; size of amber piece approximately 8.9
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<normalizedToken originalValue="×">x</normalizedToken>
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4.7
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<normalizedToken originalValue="×">x</normalizedToken>
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2.5 mm (Fig. 2), with collection label data
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<normalizedToken originalValue="“012”">"012"</normalizedToken>
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, in Andris Bukejs collection, maintained at Institute of Life Sciences and Technologies, Daugavpils University (Daugavpils, Latvia). This amber piece was found elsewhere in the area of the Curonian Spit.
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</paragraph>
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<caption pageId="3" pageNumber="104">
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<paragraph pageId="3" pageNumber="104">
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Figures 1-2.
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<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion bukejsi" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="species" species="bukejsi">Bembidion bukejsi</taxonomicName>
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sp. n., light microscopic images of the holotype. 1 dorsal aspect 2 general view of the fossil with contours of the amber piece.
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</paragraph>
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</caption>
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</subSubSection>
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<subSubSection pageId="3" pageNumber="104" type="preservation status">
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<paragraph pageId="3" pageNumber="104">Preservation status.</paragraph>
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<paragraph pageId="3" pageNumber="104">
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The amber piece and its
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<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Bembidion</taxonomicName>
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fossil are in a comparatively very good conservation state. Most parts of the piece are clear, and the beetle body is well visible using light microscopy (Figs 1, 3-5). The mouth part is partly covered by milky coating and thus mandibles and maxillae cannot be investigated by light microscopy. The body is partly covered by minute particles (granules or tiny bubbles) of unknown origin. The exoskeleton of the specimen is partly slightly shrunken and thus dissociated from the inclusion wall (Figs 6-8). Four apical tarsomeres of the left proleg are lost. The aedeagus is well preserved and could be reconstructed together with its endophallic folding structures using micro-CT (Figs 12-17, 26).
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</paragraph>
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<caption pageId="3" pageNumber="104">
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<paragraph pageId="3" pageNumber="104">
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Figures 3-5.
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<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion bukejsi" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="species" species="bukejsi">Bembidion bukejsi</taxonomicName>
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sp. n., light microscopic images of the holotype. 3 anterior portion of body, dorsal aspect 4 anterior portion of body, ventral aspect 5 left lateral aspect.
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</paragraph>
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</caption>
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<caption pageId="3" pageNumber="104">
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<paragraph pageId="3" pageNumber="104">
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Figures 6-8.
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<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion bukejsi" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="species" species="bukejsi">Bembidion bukejsi</taxonomicName>
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sp. n., volume rendering of the holotype. 6 dorsal aspect 7 left lateral aspect 8 ventral aspect.
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</paragraph>
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</caption>
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</subSubSection>
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<subSubSection pageId="3" pageNumber="104" type="syninclusions">
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<paragraph pageId="3" pageNumber="104">Syninclusions.</paragraph>
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<paragraph pageId="3" pageNumber="104">One stellate hair and numerous dirt particles.</paragraph>
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</subSubSection>
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<subSubSection pageId="3" pageNumber="104" type="description">
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<paragraph pageId="3" pageNumber="104">Description.</paragraph>
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<paragraph pageId="3" pageNumber="104">Body length: 3.5 mm.</paragraph>
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<paragraph pageId="3" pageNumber="104">Colour: The whole body surface appears blackish, very shiny, with metallic lustre; variation in colouration of the different parts of the beetle body is not recognizable.</paragraph>
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<paragraph pageId="3" pageNumber="104">
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Microsculpture: Surface of head including labrum with deeply engraved isodiametric sculpticells, pronotum with less deeply engraved slightly transverse meshes, elytral intervals with very finely engraved transverse meshes which are much smaller than on head and pronotum and which are not visible below magnification of
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<normalizedToken originalValue="×">x</normalizedToken>
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100.
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</paragraph>
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<paragraph pageId="3" pageNumber="104">
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Head: Moderately large and transverse; length 0.75 mm, width 0.80 mm. Mandibles moderately slender. Labrum with apical margin slightly concave, dorsally with three pairs of setae near apical margin. Clypeus with one pair of setae in normal position. Shape and setation of maxillary palpi as typical for
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<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Bembidion</taxonomicName>
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, with apical segment subulate, approx. 2/5 of length of penultimate segment; penultimate segment markedly broadened towards apex. Antennae rather short, with pedicellus approx. 1.5 times longer than broad, and with two antennomeres extending beyond the pronotal base. Mentum and submentum distinct, mentum with medio-apical tooth simple, shortly rounded at tip, and with one pair of setae; pits absent. Eyes large, hemispherical, protruded; tempora very small, approx. 1/20 of eyes diameter, not visible in dorsal view. Disk moderately convex, smooth apart from the prominent microsculpture. Frontal furrows very shallow, very short, absent on disk. Supraorbital furrows flat, without punctures; two supraorbital setae present and in normal position for
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<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Bembidion</taxonomicName>
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.
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</paragraph>
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<paragraph pageId="3" pageNumber="104">Prothorax: Pronotum moderately large, length 0.76 mm, width 1.04 mm, transverse (width/length = 1.37), 1.3 times broader than head, subcordate, broadest slightly before middle, with sides faintly concave in posterior third. Laterobasal angles large, almost rectangular, not protruded laterally. Basal margin 1.1 times broader than apical margin. Disk moderately convex, smooth. Anterior margin finely convex in middle, lateroapical angles distinctly protruded, rounded. Posterior margin not beaded, slightly convex in middle, slightly sinusoidal towards laterobasal angles, latter very faintly shifted anteriad with respect to posterior margin of pronotum. Median longitudinal impression deep in middle, deepest before posterior transverse impression, but absent near pronotal apex and base; anterior transverse impression very shallow, smooth; posterior transverse impression moderately deep, smooth; laterobasal foveae large and rounded, moderately deep, smooth. Lateral gutter narrow throughout, smooth. Laterobasal carina long and straight, approx. 1/3 of length of pronotum. Both lateral and laterobasal setae present, with the lateral seta located slightly before middle of pronotum. Proepisternum glabrous, smooth.</paragraph>
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<paragraph pageId="3" pageNumber="104">Pterothorax: Elytra moderately convex on disc, in dorsal view narrow ovate, length 2.05 mm, width 1.37 mm, length/width = 1.50, widest near anterior third, distinctly wider than pronotum (width of elytra/width of pronotum = 1.32). Surface and lateral border glabrous and smooth apart from the primary elytral setation. Shoulders moderately broad with humeral margin angulate: The lateral bead forms an almost right angle with the abbreviated basal bead; the latter extends to the tip of the 4th elytral stria. Crista clavicularis absent. Sides with preapical sinuation indistinct; subapical plica present. Parascutellar stria moderately long, parascutellar seta present. All striae complete, deeply impressed, impunctate, with intervals convex; apical stria (= common prolongation of the 5th, 6th, and 7th striae) deeply impressed from level of the apical cross of 5th and 6th stria towards apex; recurrent stria lacking. Ninth interval moderately broadened from level of humeral umbilicate series towards apex. Each elytron with two discal setae in third interval, with relevant pores located close to, but separated from, third stria. Preapical seta located in the deepened apical portion of the seventh stria; the fine apical seta located at apical margin. Umbilicate series consist of eight setae: four humeral setae, with distance between first and second as well as second and third setae slightly larger than that between third and fourth setae; the fourth seta is located distinctly basad of the level of the anterior discal seta; both the subapical setae are markedly advanced and located at the beginning of the apical elytral third; two apical setae of the umbilicate series situated anterior of the junction of the eighth stria and lateral gutter. Metepisternum long, glabrous and smooth, with outer margin 1.6 times longer than anterior margin. Metasternal process without borders, moderately convex in middle. Hindwings fully developed.</paragraph>
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<paragraph pageId="3" pageNumber="104">
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Abdomen: Abdominal sternites
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<normalizedToken originalValue="V–VII">V-VII</normalizedToken>
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each with one (male) pair of setae near apical margin; surfaces smooth, without hairs or micropunctures.
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</paragraph>
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<paragraph pageId="3" pageNumber="104">Legs: Relatively short, unmodified, femora moderately robust, protibiae straight and moderately dilated towards apex. First protarsomere markedly dilated, second protarsomere moderately dilated with apicolateral projection on inner margin.</paragraph>
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<paragraph pageId="3" pageNumber="104">
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Male genitalia (Figs 12-17, 26): Shape and size of median lobe as well as general structures of endophallus similar to species of the
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<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Bembidion</taxonomicName>
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subgenera
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<taxonomicName class="Insecta" family="Carabidae" genus="Bracteon" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bracteon" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Bracteon</taxonomicName>
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Bedel, 1879, and
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<taxonomicName class="Insecta" family="Carabidae" genus="Odontium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Odontium" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Odontium</taxonomicName>
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LeConte, 1848 (see
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<bibRefCitation author="Maddison, DR" journalOrPublisher="Bulletin of the Museum of Comparative Zoology" pageId="9" pageNumber="110" pagination="142 - 299" title="Systematics of the Holarctic beetle subgenus Bracteon and related Bembidion (Coleoptera: Carabidae)." volume="153" year="1993">Maddison 1993</bibRefCitation>
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: 257): Median lobe moderately large, moderately slender, in lateral view moderately bent, with terminal lamella distinct, short tongue-shaped, slightly bent ventrad. Details of the parameres could not be recovered. Endophallus (in inverted positon) with a large ostidial flag with its internal tip distinctly bent back to the dorsal side of median lobe; dorsal field of ostium distinct; dorsal plate very large with left side more than right side. Below the dorsal plate an extended folding structure is developed (central fold system) which covers the central sclerite and the brush sclerite complexes dorsally and, which proceed in the flagellum distally; latter rather short, almost straight, not extending to the dorsal tip of the ostidial flag. Central and brush sclerite complexes both moderately large and situated side by side in middle portion of median lobe, overlaying each other in lateral view; central sclerite with left lobe lenticular and with right lobe large, situated more basad; brush sclerite apico-ventrally markedly prolonged with complicate folding structures.
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</paragraph>
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<caption pageId="3" pageNumber="104">
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<paragraph pageId="3" pageNumber="104">
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Figures 9-13.
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<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion bukejsi" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="species" species="bukejsi">Bembidion bukejsi</taxonomicName>
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sp. n., holotype, volume rendering of selected body parts 9 caudal aspect of body 10 tarsomeres and distal portion of tibia of right proleg. 11 frontal section of body (ventral aspect) showing position of the abdominal segment IX which surrounds the aedeagus 12 abdominal segment IX and aedeagal median lobe, left lateral aspect 13 sagittal section of aedeagal median lobe, left lateral aspect. Abbreviations: antc = antecosta; bow-l = left wall of basal orfice; bsc-fd = folding structures originating from the endophallic brush sclerite; cfd = central folding system of endophallus; csc = central sclerite of endophallus (left lobe); dpl = dorsal plate of endophallus; mla = aedeagal median lobe apex; mlb = aedeagal median lobe base; mlw-d = dorsal wall of median lobe; mlw-v = ventral wall of median lobe; mtg IX = mediotergite IX; omp = ostial microtrichial patch; pm-l = left paramere (basal portion).
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</paragraph>
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</caption>
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<caption pageId="3" pageNumber="104">
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<paragraph pageId="3" pageNumber="104">
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Figures 14-17.
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<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion bukejsi" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="species" species="bukejsi">Bembidion bukejsi</taxonomicName>
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sp. n., holotype, visualization of endophallic structures using micro-CT. 14 volume rendering of the aedeagal median lobe with sclerotized endophallic structures highlighted in colors 15-16 parts of transverse sections through the abdomen with aedeagus (for position of slices see Fig. 14) 17 sclerotized endophallic structures separated and highlighted in colors using serial sectioning (a-d, left lateral aspect; e, right lateral aspect; f-g, dorsal aspect). Colour coding: beige = internal fold originating from the brush sclerite; green = central sclerite; khaki = dorsal field; light blue = dorsal plate; marine blue = ostidial flag; red = central fold system; white = unknown sclerite; yellow = brush sclerite. Abbreviations: bsc = brush sclerite; bsc-fd-i = internal fold originating from the brush sclerite; bsc-fd-v = ventral prolongation of the brush sclerite; cfd = central folding system; csc = central sclerite; dpl = dorsal plate; mlw = wall of median lobe; scu = unknown sclerite; st = sternite.
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</paragraph>
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</caption>
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</subSubSection>
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<subSubSection pageId="3" pageNumber="104" type="etymology">
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<paragraph pageId="3" pageNumber="104">Derivatio nominis.</paragraph>
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<paragraph pageId="3" pageNumber="104">The species epithet is a dedication to the collector Andris Bukejs, which kindly allowed us to investigate this unique specimen.</paragraph>
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</subSubSection>
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<subSubSection lastPageId="5" lastPageNumber="106" pageId="3" pageNumber="104" type="relationships and recognition">
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<paragraph pageId="3" pageNumber="104">Relationships and recognition.</paragraph>
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<paragraph pageId="3" pageNumber="104">
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The angulate humeral margin together with the position of the elytral discal setae separated from third stria and the shape of the endophallic structures provide important evidence for probable relationships of the fossil species. An angulate humeral margin is also developed in species of the
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<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Bembidion</taxonomicName>
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sensu lato lineages
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<taxonomicName class="Insecta" family="Carabidae" genus="Andrewesa" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Andrewesa" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Andrewesa</taxonomicName>
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Netolitzky, 1931,
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<taxonomicName class="Insecta" family="Carabidae" genus="Hoquedela" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hoquedela" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Hoquedela</taxonomicName>
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<normalizedToken originalValue="Müller-Motzfeld">Mueller-Motzfeld</normalizedToken>
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, 1988,
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<taxonomicName class="Insecta" family="Carabidae" genus="Peryphophila" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Peryphophila" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Peryphophila</taxonomicName>
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Netolitzky, 1939,
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<taxonomicName class="Insecta" family="Carabidae" genus="Phyla" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phyla" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Phyla</taxonomicName>
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Motschulsky, 1844,
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<taxonomicName class="Insecta" family="Carabidae" genus="Plataphodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Plataphodes" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Plataphodes</taxonomicName>
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Ganglbauer, 1891, as well as in the
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<taxonomicName class="Insecta" family="Carabidae" genus="Hydrium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hydrium" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Hydrium</taxonomicName>
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and
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<taxonomicName class="Insecta" family="Carabidae" genus="Odontium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Odontium" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Odontium</taxonomicName>
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complexes sensu
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<bibRefCitation author="Maddison, DR" journalOrPublisher="Molecular Phylogenetics and Evolution," pageId="9" pageNumber="110" pagination="533 - 576" title="Phylogeny of Bembidion and related ground beetles (Coleoptera: Carabidae: Trechinae: Bembidiini: Bembidiina)." url="http://dx.doi.org/10.1016/j.ympev.2012.01.015" volume="63" year="2012">Maddison (2012)</bibRefCitation>
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.
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<taxonomicName class="Insecta" family="Carabidae" genus="Pekinium" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Pekinium" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Pekinium</taxonomicName>
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Csiki, 1901, is another taxon described based on a species with angulate humeral margin, however, the type specimen seems lost and the state of the taxon remain questionable (
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<bibRefCitation author="Toledano, L" journalOrPublisher="Bollettino del Museo Civico di Storia Naturale di Venezia" pageId="9" pageNumber="110" pagination="97 - 114" title="Three subgenera of Bembidion new to China, with description of a new species (Coleoptera, Carabidae)." volume="48" year="1998">Toledano and Sciaky 1998</bibRefCitation>
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) and will thus not further considered here.
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</paragraph>
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<paragraph pageId="3" pageNumber="104">
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Based on a comprehensive phylogenetic analysis of
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<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Bembidion</taxonomicName>
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and related ground beetles,
|
||
<bibRefCitation author="Maddison, DR" journalOrPublisher="Molecular Phylogenetics and Evolution," pageId="9" pageNumber="110" pagination="533 - 576" title="Phylogeny of Bembidion and related ground beetles (Coleoptera: Carabidae: Trechinae: Bembidiini: Bembidiina)." url="http://dx.doi.org/10.1016/j.ympev.2012.01.015" volume="63" year="2012">Maddison (2012)</bibRefCitation>
|
||
has shown that the character state 'angulate humeral
|
||
<normalizedToken originalValue="margin’">margin'</normalizedToken>
|
||
evolved probably five times independently within
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Bembidion</taxonomicName>
|
||
sensu lato. In
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Phyla" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phyla" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Phyla</taxonomicName>
|
||
, the abbreviated basal bead extends to the tip of the 5th elytral stria and is therefore distinctly shorter than in
|
||
<taxonomicName lsidName="B. bukejsi" pageId="3" pageNumber="104" rank="species" species="bukejsi">B. bukejsi</taxonomicName>
|
||
sp. n.. The same holds true for
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Peryphophila" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Peryphophila" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Peryphophila</taxonomicName>
|
||
, which was not included in the analysis of
|
||
<bibRefCitation author="Maddison, DR" journalOrPublisher="Molecular Phylogenetics and Evolution," pageId="9" pageNumber="110" pagination="533 - 576" title="Phylogeny of Bembidion and related ground beetles (Coleoptera: Carabidae: Trechinae: Bembidiini: Bembidiina)." url="http://dx.doi.org/10.1016/j.ympev.2012.01.015" volume="63" year="2012">Maddison (2012)</bibRefCitation>
|
||
. Species of both taxa can be additionally distinguished from
|
||
<taxonomicName lsidName="B. bukejsi" pageId="3" pageNumber="104" rank="species" species="bukejsi">B. bukejsi</taxonomicName>
|
||
sp. n. in the elytral discal setae, which are situated in the third stria, and by its very different endophallic morphology. In
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Phyla" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phyla" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Phyla</taxonomicName>
|
||
, an additional endophallic sclerite is developed basad of the central sclerite complex (probably homolog to the
|
||
<normalizedToken originalValue="“N-sclerite”">"N-sclerite"</normalizedToken>
|
||
of the
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Notaphus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Notaphus" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Notaphus</taxonomicName>
|
||
series sensu
|
||
<bibRefCitation author="Maddison, DR" journalOrPublisher="Molecular Phylogenetics and Evolution," pageId="9" pageNumber="110" pagination="533 - 576" title="Phylogeny of Bembidion and related ground beetles (Coleoptera: Carabidae: Trechinae: Bembidiini: Bembidiina)." url="http://dx.doi.org/10.1016/j.ympev.2012.01.015" volume="63" year="2012">Maddison 2012</bibRefCitation>
|
||
, Fig. 19), the central sclerite is irregularly shaped, and the dorsal field is lacking. In
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Peryphophila" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Peryphophila" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Peryphophila</taxonomicName>
|
||
, the central sclerite is markedly small and the aedeagal median lobe is much more slender compared to
|
||
<taxonomicName lsidName="B. bukejsi" pageId="3" pageNumber="104" rank="species" species="bukejsi">B. bukejsi</taxonomicName>
|
||
sp. n. (Fig. 22). In the
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Plataphus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Plataphus" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Plataphus</taxonomicName>
|
||
complex sensu
|
||
<bibRefCitation author="Maddison, DR" journalOrPublisher="Molecular Phylogenetics and Evolution," pageId="9" pageNumber="110" pagination="533 - 576" title="Phylogeny of Bembidion and related ground beetles (Coleoptera: Carabidae: Trechinae: Bembidiini: Bembidiina)." url="http://dx.doi.org/10.1016/j.ympev.2012.01.015" volume="63" year="2012">Maddison (2012)</bibRefCitation>
|
||
, including
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Plataphodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Plataphodes" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Plataphodes</taxonomicName>
|
||
, the central sclerite is only hardly sclerotized or sometimes lacking and thus, the endophallic structures are very differently developed (Fig. 20) compared to the situation in
|
||
<taxonomicName lsidName="B. bukejsi" pageId="3" pageNumber="104" rank="species" species="bukejsi">B. bukejsi</taxonomicName>
|
||
sp. n.. Moreover, also in species of the
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Plataphus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Plataphus" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Plataphus</taxonomicName>
|
||
complex, the elytral discal setae are situated in the third stria. In the High Asian taxon
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Hoquedela" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hoquedela" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Hoquedela</taxonomicName>
|
||
, the elytral discal setae are separated from the third stria similarly to the organization in
|
||
<taxonomicName lsidName="B. bukejsi" pageId="3" pageNumber="104" rank="species" species="bukejsi">B. bukejsi</taxonomicName>
|
||
sp. n.. However,
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Hoquedela" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hoquedela" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Hoquedela</taxonomicName>
|
||
differs from
|
||
<taxonomicName lsidName="B. bukejsi" pageId="3" pageNumber="104" rank="species" species="bukejsi">B. bukejsi</taxonomicName>
|
||
sp. n. by the shape of the aedeagal median lobe (Fig. 18), which is markedly robust with a swollen middle portion and very stout apex, by an extensively sclerotized folding structure of endophallus particularly near median lobe ostium, and by modified mesotibia which are suggestively sinuate due to slightly convex interior surface in middle of tibia and slightly inwardly bent tibial apex.
|
||
</paragraph>
|
||
<caption pageId="3" pageNumber="104">
|
||
<paragraph pageId="3" pageNumber="104">
|
||
Figures 18-26. Aedeagal median lobes and endophallic structures of recent
|
||
<taxonomicName lsidName="Bembidiina" pageId="3" pageNumber="104" rank="subtribe" subtribe="Bembidiina">Bembidiina</taxonomicName>
|
||
(18-25) and the fossil
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion bukejsi" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="species" species="bukejsi">Bembidion bukejsi</taxonomicName>
|
||
sp. n. (26), left lateral view. 18
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Hoquedela" higherTaxonomySource="CoL" infraspecific-rank="k." kingdom="Animalia" lsidName="Hoquedela" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="subspecies" subspecies="kirschenhoferi">Hoquedela k. kirschenhoferi</taxonomicName>
|
||
<normalizedToken originalValue="Müller-Motzfeld">Mueller-Motzfeld</normalizedToken>
|
||
, 1988 19
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion (Phyla) tethys" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="species" species="tethys" subGenus="Phyla">Bembidion (Phyla) tethys</taxonomicName>
|
||
Netolitzky, 1926 20
|
||
<taxonomicName infraspecific-rank="f." lsidName="B. (Plataphus)" pageId="3" pageNumber="104" rank="subspecies" subGenus="Plataphus" subspecies="fellmanni">B. (Plataphus) f. fellmanni</taxonomicName>
|
||
Mannerheim, 1823 21
|
||
<taxonomicName lsidName="B. (Melomalus) altaicum" pageId="3" pageNumber="104" rank="species" species="altaicum" subGenus="Melomalus">B. (Melomalus) altaicum</taxonomicName>
|
||
Gebler, 1833 22
|
||
<taxonomicName lsidName="B. (Peryphophila) eurydice" pageId="3" pageNumber="104" rank="species" species="eurydice" subGenus="Peryphophila">B. (Peryphophila) eurydice</taxonomicName>
|
||
Andrewes, 1926 23
|
||
<taxonomicName lsidName="B. (Andrewesa) patris" pageId="3" pageNumber="104" rank="species" species="patris" subGenus="Andrewesa">B. (Andrewesa) patris</taxonomicName>
|
||
Schmidt, 2010 24
|
||
<taxonomicName lsidName="B. (Bracteon) lapponicum" pageId="3" pageNumber="104" rank="species" species="lapponicum" subGenus="Bracteon">B. (Bracteon) lapponicum</taxonomicName>
|
||
Zetterstedt, 1828 25
|
||
<taxonomicName lsidName="B. (Odontium) striatum" pageId="3" pageNumber="104" rank="species" species="striatum" subGenus="Odontium">B. (Odontium) striatum</taxonomicName>
|
||
Fabricius, 1792 26
|
||
<taxonomicName lsidName="B. (Eodontium) bukejsi" pageId="3" pageNumber="104" rank="species" species="bukejsi" subGenus="Eodontium">B. (Eodontium) bukejsi</taxonomicName>
|
||
subgen. n., sp. n. (Fig. 26a, volume rendering with highlighted structures, see also Fig. 14; Fig. 26b, schematic reconstruction of putative organization of endophallic structures). Scale bar: 2.2 mm. Abbreviations: bsc = brush sclerite; bsc-fd-av = apico-ventral prolongation of the brush sclerite; csc = central sclerite; csc-ll = left lobe of central sclerite; csc-rl = right lobe of central sclerite; df = dorsal field; dpl = dorsal plate; fl = flagellum of the central fold system; N = N-sclerite (
|
||
<bibRefCitation author="Maddison, DR" journalOrPublisher="Molecular Phylogenetics and Evolution," pageId="9" pageNumber="110" pagination="533 - 576" title="Phylogeny of Bembidion and related ground beetles (Coleoptera: Carabidae: Trechinae: Bembidiini: Bembidiina)." url="http://dx.doi.org/10.1016/j.ympev.2012.01.015" volume="63" year="2012">Maddison 2012</bibRefCitation>
|
||
); of = ostidial flag.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph pageId="3" pageNumber="104">
|
||
The external shape of the aedeagal median lobe and sclerotization patterns of the endophallic structures present in
|
||
<taxonomicName lsidName="B. bukejsi" pageId="3" pageNumber="104" rank="species" species="bukejsi">B. bukejsi</taxonomicName>
|
||
sp. n. resemble those of species of
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Andrewesa" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Andrewesa" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Andrewesa</taxonomicName>
|
||
(Fig. 23), the
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Hydrium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hydrium" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Hydrium</taxonomicName>
|
||
complex and
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Odontium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Odontium" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Odontium</taxonomicName>
|
||
series (Figs 24, 25,
|
||
<bibRefCitation author="Maddison, DR" journalOrPublisher="Bulletin of the Museum of Comparative Zoology" pageId="9" pageNumber="110" pagination="142 - 299" title="Systematics of the Holarctic beetle subgenus Bracteon and related Bembidion (Coleoptera: Carabidae)." volume="153" year="1993">Maddison 1993</bibRefCitation>
|
||
: Figs 112-128), as well as in
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Liocosmius" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Liocosmius" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Liocosmius</taxonomicName>
|
||
Casey, 1918 (
|
||
<bibRefCitation author="Maddison, DR" journalOrPublisher="Zoological Journal of the Linnean Society" pageId="9" pageNumber="110" pagination="741 - 770" title="Species delimitation in the ground beetle subgenus Liocosmius (Coleoptera: Carabidae: Bembidion), including standard and next-generation sequencing of museum specimens." url="https://doi.org/10.1111/zoj.12188" volume="172" year="2014">Maddison and Cooper 2014</bibRefCitation>
|
||
: Fig. 12),
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Melomalus" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Melomalus" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Melomalus</taxonomicName>
|
||
Casey, 1918 (Fig. 21),
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Trechonepha" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Trechonepha" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Trechonepha</taxonomicName>
|
||
Casey, 1918,
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Trichoplataphus" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Trichoplataphus" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Trichoplataphus</taxonomicName>
|
||
Netolitzky, 1914 (
|
||
<bibRefCitation author="Toledano, L" journalOrPublisher="Entomologische Blaetter" pageId="9" pageNumber="110" pagination="371 - 406" title="Revision of the Bembidionkara Andrewes, 1921 species group and notes on the Palaearctic species of Bembidion subgenus Trichoplataphus Netolitzky, 1914 (Coleoptera, Carabidae, Bembidiini)." volume="106" year="2010">Toledano and Schmidt 2010</bibRefCitation>
|
||
: Figs 33-42), and several species of the
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Ocydromus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Ocydromus" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Ocydromus</taxonomicName>
|
||
series sensu
|
||
<bibRefCitation author="Maddison, DR" journalOrPublisher="Molecular Phylogenetics and Evolution," pageId="9" pageNumber="110" pagination="533 - 576" title="Phylogeny of Bembidion and related ground beetles (Coleoptera: Carabidae: Trechinae: Bembidiini: Bembidiina)." url="http://dx.doi.org/10.1016/j.ympev.2012.01.015" volume="63" year="2012">Maddison (2012)</bibRefCitation>
|
||
. The common male genital characters of most species of these lineages are: (i) median lobe rather slender, slightly bent throughout, with (ii) terminal lamella short tongue-like, not bent backwards; (iii) endophallic central sclerite and brush sclerite complexes both moderately large and (in inverted position) situated side by side in middle portion of median lobe, overlaying each other in lateral view; (iv) central sclerite with left lobe lenticular and (v) with right lobe large and situated more basad; (vi) brush sclerite apico-ventrally markedly prolonged; (vii) dorsal plate largely developed and (viii) without additional sclerites near basal opening of median lobe. The multi-gene analyses presented by
|
||
<bibRefCitation author="Maddison, DR" journalOrPublisher="Molecular Phylogenetics and Evolution," pageId="9" pageNumber="110" pagination="533 - 576" title="Phylogeny of Bembidion and related ground beetles (Coleoptera: Carabidae: Trechinae: Bembidiini: Bembidiina)." url="http://dx.doi.org/10.1016/j.ympev.2012.01.015" volume="63" year="2012">Maddison (2012)</bibRefCitation>
|
||
support a monophyly of all these groups together with the
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Plataphus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Plataphus" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Plataphus</taxonomicName>
|
||
complex (in the following called the "
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Odontium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Odontium" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Odontium</taxonomicName>
|
||
-
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Plataphus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Plataphus" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Plataphus</taxonomicName>
|
||
-
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Ocydromus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Ocydromus" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Ocydromus</taxonomicName>
|
||
(OPO) clade). Within this highly diverse clade, a natural group '
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Ocydromus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Ocydromus" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Ocydromus</taxonomicName>
|
||
series plus
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Plataphus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Plataphus" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Plataphus</taxonomicName>
|
||
complex plus
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Trichoplataphus" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Trichoplataphus" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Trichoplataphus</taxonomicName>
|
||
' is recovered, however, the relationships of the other main lineages remain unresolved by the molecular data. In three species groups of the OPO clade, the character states 'angulate humeral
|
||
<normalizedToken originalValue="margin’">margin'</normalizedToken>
|
||
and 'elytral discal setae are separated from the third
|
||
<normalizedToken originalValue="stria’">stria'</normalizedToken>
|
||
resemble the organization observed in the fossil
|
||
<taxonomicName lsidName="B. bukejsi" pageId="3" pageNumber="104" rank="species" species="bukejsi">B. bukejsi</taxonomicName>
|
||
sp. n.,
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Andrewesa" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Andrewesa" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Andrewesa</taxonomicName>
|
||
, the
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Hydrium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hydrium" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Hydrium</taxonomicName>
|
||
and the
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Odontium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Odontium" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Odontium</taxonomicName>
|
||
complexes.
|
||
</paragraph>
|
||
<paragraph pageId="3" pageNumber="104">
|
||
Molecular data suggest that
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Andrewesa" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Andrewesa" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Andrewesa</taxonomicName>
|
||
is the sister group of the
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Hydrium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hydrium" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Hydrium</taxonomicName>
|
||
complex, while the
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Odontium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Odontium" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Odontium</taxonomicName>
|
||
complex is the sister group of the
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Hydriomicrus" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Hydriomicrus" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Hydriomicrus</taxonomicName>
|
||
complex; both the latter taxa together form the
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Odontium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Odontium" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Odontium</taxonomicName>
|
||
series (
|
||
<bibRefCitation author="Maddison, DR" journalOrPublisher="Molecular Phylogenetics and Evolution," pageId="9" pageNumber="110" pagination="533 - 576" title="Phylogeny of Bembidion and related ground beetles (Coleoptera: Carabidae: Trechinae: Bembidiini: Bembidiina)." url="http://dx.doi.org/10.1016/j.ympev.2012.01.015" volume="63" year="2012">Maddison 2012</bibRefCitation>
|
||
). The
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Hydriomicrus" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Hydriomicrus" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Hydriomicrus</taxonomicName>
|
||
complex includes species with rounded humeral margin and elytral discal setae situated in the third stria. Close relationships of the
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Hydrium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hydrium" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Hydrium</taxonomicName>
|
||
complex plus
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Andrewesa" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Andrewesa" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Andrewesa</taxonomicName>
|
||
with the
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Odontium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Odontium" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Odontium</taxonomicName>
|
||
series, are not supported by the molecular data. Thus, it remains unclear whether the character states 'angulate humeral
|
||
<normalizedToken originalValue="margin’">margin'</normalizedToken>
|
||
and 'elytral discal setae separated from third
|
||
<normalizedToken originalValue="stria’">stria'</normalizedToken>
|
||
were evolved in the OPO clade only once or even twice; in latter case, an independent evolution of these features in the
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Hydrium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hydrium" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Hydrium</taxonomicName>
|
||
complex plus
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Andrewesa" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Andrewesa" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Andrewesa</taxonomicName>
|
||
, and in the
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Odontium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Odontium" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Odontium</taxonomicName>
|
||
complex of the
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Odontium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Odontium" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Odontium</taxonomicName>
|
||
series has to be assumed.
|
||
</paragraph>
|
||
<paragraph lastPageId="4" lastPageNumber="105" pageId="3" pageNumber="104">
|
||
We could not find additional derived characters justifying a further assignment of the fossil species
|
||
<taxonomicName lsidName="B. bukejsi" pageId="3" pageNumber="104" rank="species" species="bukejsi">B. bukejsi</taxonomicName>
|
||
sp. n. with one of these three lineages of the OPO clade. Similarities with representatives of extant species groups are considered to be symplesiomorphies, e.g., the shape of the pronotum and the development of the elytral striation. In
|
||
<taxonomicName lsidName="B. bukejsi" pageId="3" pageNumber="104" rank="species" species="bukejsi">B. bukejsi</taxonomicName>
|
||
sp. n., the pronotal basolateral foveae are large and rounded as in
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Andrewesa" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Andrewesa" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Andrewesa</taxonomicName>
|
||
. In many species of the
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Hydrium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hydrium" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Hydrium</taxonomicName>
|
||
complex and in all species of the
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Odontium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Odontium" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Odontium</taxonomicName>
|
||
series, the basolateral foveae are linear impressed, and the area between foveae and laterobasal carina is markedly wide and convex. The latter represent apomorphic states. The elytral striae are deeply impressed throughout in
|
||
<taxonomicName lsidName="B. bukejsi" pageId="3" pageNumber="104" rank="species" species="bukejsi">B. bukejsi</taxonomicName>
|
||
sp. n., and likewise in
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Hydriomicrus" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Hydriomicrus" order="Coleoptera" pageId="4" pageNumber="105" phylum="Arthropoda" rank="genus">
|
||
<pageBreakToken pageId="4" pageNumber="105" start="start">Hydriomicrus</pageBreakToken>
|
||
</taxonomicName>
|
||
Casey, 1918,
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Hirmoplataphus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hirmoplataphus" order="Coleoptera" pageId="4" pageNumber="105" phylum="Arthropoda" rank="genus">Hirmoplataphus</taxonomicName>
|
||
Lindroth, 1963, and
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Pseudoperyphus" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Pseudoperyphus" order="Coleoptera" pageId="4" pageNumber="105" phylum="Arthropoda" rank="genus">Pseudoperyphus</taxonomicName>
|
||
of the
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Odontium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Odontium" order="Coleoptera" pageId="4" pageNumber="105" phylum="Arthropoda" rank="genus">Odontium</taxonomicName>
|
||
series. In all the remaining lineages of the
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Odontium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Odontium" order="Coleoptera" pageId="4" pageNumber="105" phylum="Arthropoda" rank="genus">Odontium</taxonomicName>
|
||
series as well as in
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Andrewesa" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Andrewesa" order="Coleoptera" pageId="4" pageNumber="105" phylum="Arthropoda" rank="genus">Andrewesa</taxonomicName>
|
||
and the
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Hydrium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hydrium" order="Coleoptera" pageId="4" pageNumber="105" phylum="Arthropoda" rank="genus">Hydrium</taxonomicName>
|
||
complex, the external elytral striae are more or less distinctly more shallowly impressed before the apex. Latter is considered an apomorphic character state.
|
||
</paragraph>
|
||
<paragraph lastPageId="5" lastPageNumber="106" pageId="4" pageNumber="105">
|
||
Due to the impunctate elytral stria
|
||
<taxonomicName lsidName="B. bukejsi" pageId="4" pageNumber="105" rank="species" species="bukejsi">B. bukejsi</taxonomicName>
|
||
sp. n. differs strikingly from all species of
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Andrewesa" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Andrewesa" order="Coleoptera" pageId="4" pageNumber="105" phylum="Arthropoda" rank="genus">Andrewesa</taxonomicName>
|
||
, the
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Hydrium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hydrium" order="Coleoptera" pageId="4" pageNumber="105" phylum="Arthropoda" rank="genus">Hydrium</taxonomicName>
|
||
complex and the
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Odontium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Odontium" order="Coleoptera" pageId="4" pageNumber="105" phylum="Arthropoda" rank="genus">Odontium</taxonomicName>
|
||
series. However, impunc
|
||
<pageBreakToken pageId="5" pageNumber="106" start="start">tate</pageBreakToken>
|
||
or indistinctly punctate elytral striae are also present in other lineages of the clade, e.g.,
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Melomalus" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Melomalus" order="Coleoptera" pageId="5" pageNumber="106" phylum="Arthropoda" rank="genus">Melomalus</taxonomicName>
|
||
, the
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Ocydromus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Ocydromus" order="Coleoptera" pageId="5" pageNumber="106" phylum="Arthropoda" rank="genus">Ocydromus</taxonomicName>
|
||
series, and the
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Plataphus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Plataphus" order="Coleoptera" pageId="5" pageNumber="106" phylum="Arthropoda" rank="genus">Plataphus</taxonomicName>
|
||
complex. This character shows also continuous variation in other
|
||
<taxonomicName lsidName="Bembidiina" pageId="5" pageNumber="106" rank="subtribe" subtribe="Bembidiina">Bembidiina</taxonomicName>
|
||
clades and is therefore not informative with regard to its phylogenetic implications.
|
||
</paragraph>
|
||
<paragraph pageId="5" pageNumber="106">
|
||
Due to the peculiar combination of the character states observed in
|
||
<taxonomicName lsidName="B. bukejsi" pageId="5" pageNumber="106" rank="species" species="bukejsi">B. bukejsi</taxonomicName>
|
||
sp. n., we conclude that this fossil species represents an extinct lineage of the OPO clade, probably related to either
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Andrewesa" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Andrewesa" order="Coleoptera" pageId="5" pageNumber="106" phylum="Arthropoda" rank="genus">Andrewesa</taxonomicName>
|
||
plus the
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Hydrium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hydrium" order="Coleoptera" pageId="5" pageNumber="106" phylum="Arthropoda" rank="genus">Hydrium</taxonomicName>
|
||
complex or the
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Odontium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Odontium" order="Coleoptera" pageId="5" pageNumber="106" phylum="Arthropoda" rank="genus">Odontium</taxonomicName>
|
||
series, or to both of these sub-clades. Given the present state of knowledge, the assignment to one of the known subgeneric taxa is impossible and therefore we decided to describe a new subgenus for this fossil representative of
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion" order="Coleoptera" pageId="5" pageNumber="106" phylum="Arthropoda" rank="genus">Bembidion</taxonomicName>
|
||
(see below).
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection pageId="5" pageNumber="106" type="remarks">
|
||
<paragraph pageId="5" pageNumber="106">Remarks on biogeography and ecology.</paragraph>
|
||
<paragraph pageId="5" pageNumber="106">
|
||
The above mentioned OPO clade represents one of the most species rich clades of
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion" order="Coleoptera" pageId="5" pageNumber="106" phylum="Arthropoda" rank="genus">Bembidion</taxonomicName>
|
||
sensu lato (
|
||
<bibRefCitation author="Maddison, DR" journalOrPublisher="Molecular Phylogenetics and Evolution," pageId="9" pageNumber="110" pagination="533 - 576" title="Phylogeny of Bembidion and related ground beetles (Coleoptera: Carabidae: Trechinae: Bembidiini: Bembidiina)." url="http://dx.doi.org/10.1016/j.ympev.2012.01.015" volume="63" year="2012">Maddison 2012</bibRefCitation>
|
||
) with hundreds of recent species occurring mainly in the Holarctic region and, in much smaller number, also in the Oriental region and Africa. Species of the OPO clade are adapted to very different climates (tropical: e.g.,
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Microserrullula" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Microserrullula" order="Coleoptera" pageId="5" pageNumber="106" phylum="Arthropoda" rank="genus">Microserrullula</taxonomicName>
|
||
, to arctic: e.g., some species of
|
||
<taxonomicName class="Insecta" family="Carabidae" genus="Plataphus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Plataphus" order="Coleoptera" pageId="5" pageNumber="106" phylum="Arthropoda" rank="genus">Plataphus</taxonomicName>
|
||
), and prefer very different habitats although most of them are ripicolous or paludicolous. The occurrence of
|
||
<taxonomicName lsidName="B. bukejsi" pageId="5" pageNumber="106" rank="species" species="bukejsi">B. bukejsi</taxonomicName>
|
||
sp. n. in the Baltic amber forest as the first fossil representative of the OPO clade is therefore in accord with the expected distribution of that clade during the Early Cenozoic.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |